环境条件对植物稳定碳同位素组成的影响

植物稳定碳同位素技术是近年兴起的一项快速、可靠的技术。利用稳定碳同位素技术可以揭示碳同化的过程的许多方面的信息。13C和12C同位素效应,使它们在进行碳循环时发生稳定碳同位素的分馏。植物光合作用过程中CO2经气孔扩散分差和RUBPCase及PEPCase羧化分馏是造成植物稳定碳同位素比率(R=13C/12C)不同于源CO2中碳同位素比率的主要原因。遗传因素和环境因子同时决定植物碳同位素组成。植物稳定碳同位素技术同时还是古气候重建和预测未来环境变化的理论基础。本文综述了光照、温度、水分、二氧化碳、矿质营养、盐分和大气污染物等环境因素对植物稳定碳同位素组成影响方面的研究进展。     

中国暖温带落叶阔叶林中某些树种的13C自然丰度:δ13C值及其生态学意义

利用质谱分析仪对暖温带地区落叶阔叶林优势种稳定碳同位素的分析发现不同树种叶片的稳定碳同位素比率差别较大 ,大多数优势种叶片δ13C值在 -2 4.75 1‰± 0 .85 4‰～ -2 8.1 1 3‰± 1 .5 1 9‰之间。叶片的δ13C值可以分为 3个等级 , 级 ,叶片的 δ13C≥ -2 5 .5‰ , 级 ,叶片的 δ13C值在 -2 5 .5‰～ -2 7.5‰之间 , 级 ,叶片的 δ13C≤ -2 7.5‰ ,由于δ13C值在一定程度上能够反映植物的生理生态特性 ,这表明所研究的植物在生理生态特性方面也可以分为 3个类型。同时 ,由于植物的不同器官具有不同的生理生态特性 ,导致器官对 13C具有不同的分馏特性 ,也导致器官之间的δ13C值产生差异 ,分析结果显示树干、根和小枝的δ13C值一般要较叶片的δ13C值高 ,但不同树种又各具特点。生境的差异是影响稳定碳同位素比率的另一个重要原因 ,良好生境条件下生长的植物的δ13C值一般较生长在干旱瘠薄生境下的低。     

树木叶片稳定碳同位素分馏对环境梯度的响应

除了自身的遗传因素外,植物对稳定碳同位素的分馏还受温度、水分状况、光照等多种环境因素的影响。通过对生长在中国北热带、亚热带和温带的13个气候区树木的稳定碳同位素分馏与环境要素的关系的研究得出,不同气候条件下树木叶片稳定碳同位素比率(δ13C)和稳定碳同位素分辨率(Δ)差异明显,分别在-23.759‰--33.914‰和14.581‰-24.354‰的范围内。且由南向北,δ13C值呈现逐渐加重的趋势、而Δ值逐渐减小。年均温度、相对湿度、降雨量及无霜期等与δ13C值呈显著负相关、与Δ值显著正相关;年均日照时数、气温年较差、蒸发量、纬度及海拔等与δ13C值呈显著正相关、而与Δ值显著负相关;而经度、年极端最高气温等与δ13C和Δ值的相关性较弱。δ13C和Δ值的变化幅度和趋势在年均温度、年较差、无霜期、活动积温分别达到约19℃、23℃、309d、6000℃出现转折,这些指标在此基础上继续增加反而使叶片δ13C值增大、Δ值减小。而当年均降雨量和相对湿度分别超过1105mm、75%以及蒸发量小于约1700mm后,δ13C和Δ值随之变化的幅度也变小。结果表明在亚热带和温带的分界线以南,水热条件均能满足植物正常生理活动的需要,环境条件的相对改变对植物生理活动及叶片稳定碳同位素分馏的影响较小;而在北方,干旱、低温等极端的外界环境条件加剧了对植物生长和生理指标的影响、造成了不同环境梯度下叶片稳定碳同位素分馏的差异。     

δ13C在植物生态学研究中的应用

稳定碳同位素技术已成为研究植物与环境之间关系最有效的方法之一。由于植物羧化效率的不同、12 C和13 C在植物体内迁移速率以及外界环境的不同,不同植物体内稳定性碳同位素比率(δ13 C值)有一定的差异。该文概述了稳定碳同位素的基本理论,并从气孔导度、叶肉细胞导度、叶片羧化效率分析了δ13 C变化的生物学机理;对近年来国内外有关不同环境因子对植物δ13 C值的影响、δ13 C值在群落及生态系统水平(以功能群、群落冠层及树轮为重点)、以及δ13 C值在碳循环中的应用研究进展进行综述,为以后稳定碳同位素研究提供参考。     

碳稳定同位素技术在植物水分胁迫研究中的应用

植物体的碳稳定同位素组成主要由植物本身的生物学特性决定 ,但环境胁迫对其影响也十分明显。综述了碳稳定同位素技术在研究植物水分利用效率、生物量高低及判断历史气候依据等研究领域的进展 ,阐明了植物体的 δ1 3C值对干旱、盐分及其它环境因素的变化所引起的水分胁迫的响应 ,并对碳稳定同位素对水分胁迫的响应机理进行了归纳和推断     

哺乳动物化石牙齿釉质的碳、氧同位素组成与古气候重建方面的研究进展

哺乳动物化石牙齿釉质能有效地抵抗成岩作用的影响,并保存原始的碳、氧稳定同位素组成信息。通过对食草类哺乳动物化石牙齿釉质同位素组成的分析可以重建这些动物生存时代的古气候。釉质碳酸盐的碳同位素组成响应于取食植物的碳同位素组成,藉此可以恢复陆地生态系统中C3和C4植物的分布比例,这个比例主要受控于大气的CO2水平、气压和光照等因素,与不同的纬度带和气候带有关;釉质磷酸盐的氧同位素组成响应于饮用水的氧同位素组成,饮用水来源于大气降水,其氧同位素组成响应于温度变化,所以釉质的氧同位素组成与气温之间有显著的线性相关,藉此可以计算地史时期的年平均温度、绘制气候的冷暖变化曲线。     

青藏高原东部高山植物的水分利用效率与氮素利用效率研究

对生长在青藏高原东部隶属于23科、49属的71种高山植物(包括多年生和一年生植物)的稳定碳同位素比值、氮含量以及碳/氮比率进行了分析,并以稳定碳同位素比值及碳/氮比率来分别指示植物的水分利用效率和氮素利用效率.结果表明:(1)多年生植物稳定碳同位素比值显著高于一生年植物,而碳/氮比率显著低于一年生植物(P<0.01),氮含量两者无显著性差异.(2)多年生植物和一年生植物的稳定碳同位素比值均与碳/氮比率呈显著负相关(-0.643**和-0.707),而与氮含量均无明显相关性.研究证实,在自然条件下多年生植物的水分利用效率比一年生植物更高,而氮素利用效率却更低;高山植物水分利用效率和氮素利用效率存在明显的权衡",即植物不能同时提高水分利用效率和氮素利用效率,高水分利用效率的代价是降低氮素利用效率,青藏高原不同植物即使在相同环境条件下具有不同适应对策.     

浮游植物稳定碳同位素分馏机制及环境应用

浮游植物驱动水体碳循环,是解释全球碳生物地球化学过程的关键因素。相对于陆地植物,浮游植物碳在合成过程中的分馏机制较为复杂。为了系统了解藻类碳同位素分馏过程及其同位素在环境中的应用,本文调研了国内外的相关研究进展,并以蓝藻、绿藻和硅藻为代表,阐述了浮游植物碳浓缩及其同位素分馏的机制。此外,本文还总结了目前藻类碳同位素信号的变化范围及其影响因素,并探讨了藻类碳同位素在古海洋、水生食物网与水体碳循环研究中的应用。     

控温条件下C3、C4草本植物碳同位素组成对温度的响应

采取人工控制实验,探讨了6种C3、C4草本植物在昼/夜温度指标为20/12℃!36/28℃的范围内植物碳同位素组成(δ13C)及其对温度变化的响应,并结合植物比叶面积(SLA)、胞间CO2浓度(ci)与环境CO2浓度(ca)的比值、碳同化率(净光合速率Pn/胞间CO2浓度ci)等光合生长指标对植物δ13C的影响进行了分析。结果表明:所有C3、C4植物样品的δ13C值分别变化在-28.3‰!-32.1‰和-14.4‰!-17.6‰之间;在C3植物中,油菜δ13C值分布范围最集中,位于-31.1‰!-32.1‰之间;C4植物中,谷子δ13C值分布范围最窄。在控制的温度范围内,3种C3植物的平均δ13C值随温度升高而显著变低,而C4植物δ13C平均值与温度呈先增大后减小的抛物型关系,但线性回归结果未达到显著水平(P0.05)。单个植物种的δ13C值对温度的响应不同,茄子、高粱的δ13C值与温度呈线性负相关,其它4种植物与温度均呈二次抛物线关系,这可能与不同植物种具有不同的光合最适温度以及植物δ13C分馏对温度变化的敏感程度不同有关。     

高寒草甸消费者种群稳定碳、氮同位素组成的海拔分异

通过测定青藏高原东部高寒区不同海拔主要植物和消费者种群（雀形目鸟类和小型哺乳类）的稳定碳、氮同位素比值,研究了高寒草甸消费者种群同位素组成特征及其影响因素。结果表明：植物平均稳定碳同位素随海拔升高表现出明显的增加趋势;消费者种群稳定碳、氮同位素随海拔的升高有明显的富集效应。雀形目鸟类种群稳定碳、氮同位素随海拔的升高均呈明显增加趋势;小型哺乳类稳定碳同位素随海拔上升不明显,而稳定氮同位素具有明显的增大趋势。消费者种群稳定碳、氮同位素沿海拔梯度的富集效应,一方面与植物稳定同位素在海拔梯度上的富集密切相关;另一方面与海拔和纬度变化所引起一系列环境因子的变化在一定程度上影响到动物的稳定同位素分布模式。另外,在高寒草甸地区,与雀形目鸟类相比,小型哺乳类在动物稳定同位素组成的代谢过程中更容易受到环境改变的影响。     

Factors affecting the isotopic composition of organic matter. (1) Carbon isotopic composition of terrestrial plant materials

The stable isotope composition of the light elements (i.e., H, C, N, O and S) of organic samples varies significantly and, for C, is also unique and distinct from that of inorganic carbon. This is the result of (1) the isotope composition of reactants, (2) the nature of the reactions leading to formation and post-formational modification of the samples, (3) the environmental conditions under which the reactions took place, and (4) the relative concentration of the reactants compared to that of the products (i.e., [products]/[reactants] ratio). This article will examine the carbon isotope composition of terrestrial plant materials and its relationship with the above factors. delta13C(PDB) values of terrestrial plants range approximately from -8 to -38%, inclusive of C3-plants (-22 to -38%), C4-plants (-8 to -15%) and CAM-plants (-13 to -30%). Thus, the delta13C(PDB) values largely reflect the photosynthesis pathways of a plant as well as the genetics (i.e., species difference), delta13C(PDB) values of source CO2, relevant humidity, CO2/O2 ratios, wind and light intensity etc. Significant variations in these values also exist among different tissues, different portions of a tissue and different compounds. This is mainly a consequence of metabolic reactions. Animals mainly inherit the delta13C(PDB) values of the foods they consume; therefore, their delta13C(PDB) values are similar. The delta13C(PDB) values of plant materials, thus, contain information regarding the inner workings of the plants, the environmental conditions under which they grow, the delta13C(PDB) values of CO2 sources etc., and are unique. Furthermore, this uniqueness is passed on to their derivative matter, such as animals, humus etc. Hence, they are very powerful tools in many areas of research, including the ecological and environmental sciences.     

马尾松和杉木树干韧皮部水溶性糖 δ13C值对气象因子的响应

通过测定亚热带马尾松和杉木树干韧皮部水溶性糖δ¹³C值的连日变化,及其对天气变化过程的响应,研究δ¹³C值对短期天气变化动态的响应特征。结果显示,春季马尾松和杉木树干韧皮部水溶性糖δ¹³C日均值分别介于-26.81‰到-26.49‰之间,以及-29.26‰到-27.47‰之间,平均值分别为(-26.58±0.12)‰和(-28.67±0.65)‰。进一步分析表明,马尾松树干韧皮部水溶性糖δ¹³C值与取样之前第4天的太阳辐射、水气压亏缺、相对湿度和空气温度显著相关(P≤0.05),杉木树干韧皮部水溶性糖δ¹³C值取样之前第3天的太阳辐射、水气压亏缺和相对湿度显著相关(P≤0.05),但与空气温度的相关性不显著(P≤0.05)。在所测定的环境因子中,太阳辐射是影响马尾松和杉木树干韧皮部水溶性糖δ¹³C值的首要因素。当天降水事件可能导致马尾松和杉木树干韧皮部水溶性糖δ¹³C值连日变化出现异常波动。马尾松和杉木韧皮部水溶性糖δ¹³C值可以敏感记录短期天气变化动态。     

Carbonic anhydrase and its influence on carbon isotope discrimination during C4 photosynthesis. Insights from antisense RNA in Flaveria bidentis

In C4 plants, carbonic anhydrase (CA) facilitates both the chemical and isotopic equilibration of atmospheric CO2 and bicarbonate (HCO3-) in the mesophyll cytoplasm. The CA-catalyzed reaction is essential for C4 photosynthesis, and the model of carbon isotope discrimination (Delta13C) in C4 plants predicts that changes in CA activity will influence Delta13C. However, experimentally, the influence of CA on Delta13C has not been demonstrated in C4 plants. Here, we compared measurements of Delta13C during C4 photosynthesis in Flaveria bidentis wild-type plants with F. bidentis plants with reduced levels of CA due to the expression of antisense constructs targeted to a putative mesophyll cytosolic CA. Plants with reduced CA activity had greater Delta13C, which was also evident in the leaf dry matter carbon isotope composition (delta13C). Contrary to the isotope measurements, photosynthetic rates were not affected until CA activity was less than 20% of wild type. Measurements of Delta13C, delta13C of leaf dry matter, and rates of net CO2 assimilation were all dramatically altered when CA activity was less than 5% of wild type. CA activity in wild-type F. bidentis is sufficient to maintain net CO2 assimilation; however, reducing leaf CA activity has a relatively large influence on Delta13C, often without changes in net CO2 assimilation. Our data indicate that the extent of CA activity in C4 leaves needs to be taken into account when using Delta13C and/or delta13C to model the response of C4 photosynthesis to changing environmental conditions.     

典型滨岸草地生态系统碳稳定同位素组成及分布特征

通过测定上海市青浦区东风港百慕大、白花三叶草、高羊茅和白茅等4种典型滨岸草本植物各组织以及不同垂直深度土壤有机质δ¹³C值,对滨岸草地生态系统的植物-土壤碳稳定同位素特征进行了分析.结果表明: 白花三叶草、高羊茅属于C3植物,百慕大、白茅属于C4植物,其茎叶、凋落物和根系各组织间δ¹³C值无显著差异.C3和C4植物样带表层土壤有机质δ¹³C值随着土壤深度递增而呈现截然不同的变化特征,这与样带本底δ¹³C值以及碳稳定同位素分馏效应有关,同时还受植物根系分布深度的影响.植物输入是土壤有机碳(SOC)的最主要来源,植物有机体δ¹³C组成对土壤有机质δ¹³C值有直接影响,植物各组分δ¹³C值与土壤有机质δ¹³C值均存在极显著相关.4种草本植物样带SOC含量与δ¹³C值均呈极显著相关,其中,C3植物样带SOC含量与δ¹³C值呈线性负相关,C4植物样带SOC含量与δ¹³C值呈线性正相关.     

Woody tissue photosynthesis and its contribution to trunk growth and bud development in young plants

Stem photosynthesis can contribute significantly to woody plant carbon balance, particularly in times when leaves are absent or in ‘open’ crowns with sufficient light penetration. We explored the significance of woody tissue (stem) photosynthesis for the carbon income in three California native plant species via measurements of chlorophyll concentrations, radial stem growth, bud biomass and stable carbon isotope composition of sugars in different plant organs. Young plants of Prunus ilicifolia, Umbellularia californica and Arctostaphylos manzanita were measured and subjected to manipulations at two levels: trunk light exclusion (100 and 50%) and complete defoliation. We found that long‐term light exclusion resulted in a reduction in chlorophyll concentration and radial growth, demonstrating that trunk assimilates contributed to trunk carbon income. In addition, bud biomass was lower in covered plants compared to uncovered plants. Excluding 100% of the ambient light from trunks on defoliated plants led to an enrichment in ¹³C of trunk phloem sugars. We attributed this effect to a reduction in photosynthetic carbon isotope discrimination against ¹³C that in turn resulted in an enrichment in ¹³C of bud sugars. Taken together our results reveal that stem photosynthesis contributes to the total carbon income of all species including the buds in defoliated plants.     

高山植物叶片δ13C的海拔响应及其机理

植物 1 3C的分辨研究已成为植物生态学和全球碳循环研究的核心问题之一。植物 1 3C的分辨是环境和生物因子共同作用的综合结果 ,海拔梯度变化不仅可以造成植物生存环境的变化 ,而且还可以造成植物形态和生理特征的变化 ,因此 ,高山植物 1 3C分辨随海拔的变化为深入揭示植物 1 3C分辨的环境和生物因子的作用机理提供了非常理想的研究条件。在简单介绍植物 1 3C分辨基本理论的基础上 ,对目前国际上高山植物 1 3C分辨的海拔响应研究进行了述评。重点介绍了随海拔变化的大气 1 3C组成、温度、气压、水分等环境因子和植物叶片的气孔导度、羧化效率、氮含量和叶肉细胞导度等生物因子对高山 C3植物 1 3C分辨的影响 ,指出高山植物 1 3C分辨的海拔响应机理仍存在一些不确定性 ,为国内相关研究的开展提供了一定参考     

Fractionation of Carbon Isotopes during Biogenesis of Atmospheric Isoprene

The stable carbon isotope composition of isoprene emitted from leaves of red oak (Quercus rubra L.) was measured. Isoprene was depleted in ¹³C relative to carbon recently fixed by photosynthesis. The difference in isotope composition between recently fixed carbon and emitted isoprene was independent of the isotopic composition of the source CO₂. β-Carotene, an isoprenoid plant constituent, was depleted in ¹³C relative to whole leaf carbon to the same degree as isoprene, but fatty acids were more depleted. Isoprene emitted from leaves fed abscisic acid was much less depleted in ¹³C than was isoprene emitted from unstressed leaves. We conclude that isoprene is made from an isoprenoid precursor that is derived from acetyl-CoA made from recent photosynthate. The carbon isotope composition of isoprene in the atmosphere is likely to be slightly more negative (less ¹³C) than C₃ plant material but when plants are stressed the isotopic composition could vary.     

Carbon isotope analysis of acetaldehyde emitted from leaves following mechanical stress and anoxia

Although the emission of acetaldehyde from plants into the atmosphere following biotic and abiotic stresses may significantly impact air quality and climate, its metabolic origin(s) remains uncertain. We investigated the pathway(s) responsible for the production of acetaldehyde in plants by studying variations in the stable carbon isotope composition of acetaldehyde emitted during leaf anoxia or following mechanical stress. Under an anoxic environment, C3 leaves produced acetaldehyde during ethanolic fermentation with a similar carbon isotopic composition to C3 bulk biomass. In contrast, the initial emission burst following mechanical wounding was 5–12‰ more depleted in ¹³C than emissions under anoxia. Due to a large kinetic isotope effect during pyruvate decarboxylation catalysed by pyruvate dehydrogenase, acetyl-CoA and its biosynthetic products such as fatty acids are also depleted in ¹³C relative to bulk biomass. It is well known that leaf wounding stimulates the release of large quantities of fatty acids from membranes, as well as the accumulation of reactive oxygen species (ROS). We suggest that, following leaf wounding, acetaldehyde depleted in ¹³C is produced from fatty acid peroxidation reactions initiated by the accumulation of ROS. However, a variety of other pathways could also explain our results, including the conversion of acetyl-CoA to acetaldehyde by the esterase activity of aldehyde dehydrogenase.     

Carbon isotope composition of C4 grasses is influenced by light and water supply

The carbon isotope composition of C⁴ grasses has the potential to be used as an indicator of changes in the isotopic composition and concentration of atmospheric CO2, especially for climate reconstruction. The usefulness of C₄ grasses for this purpose hinges on the assumption that their photosynthetic discrimination against ¹³C remains constant in a wide range of environmental conditions. We tested this assumption by examining the effects of light and water stress on the carbon isotope composition of C₄ grasses using different biochemical subtypes (NADP-ME, NAD-ME, PCK) in glasshouse experiments. We grew 14 different C₄ grass species in four treatments: sun-watered, sun-drought, shade-watered and shade-drought. Carbon isotope discrimination (Δ) rarely remained constant. In general, Δ values were lowest in sun-watered grasses, greater for sun-drought plants and even higher for plants of the shade-watered treatment. The highest Δ values were generally found in the most stressed grasses, the shade-drought plants. Grasses of the NADP-ME subtype were the least influenced by a change in environmental variables, followed by PCK and NAD-ME subtypes. Water availability affected the carbon isotope discrimination less than light limitation in PCK and NAD-ME subtypes, but similarly in NADP-ME subtypes. In another experiment, we studied the effect of increasing light levels (150 to 1500 μmol photons m^?2 s^?1) on the Δ values of 18 well-watered C₄ grass species. Carbon isotope discrimination remained constant until photon flux density (PFD) was less than 700 μmol photons m^?2 s^?1. Below this light level, Δ values increased with decreasing irradiance for all biochemical subtypes. The change in A was less pronounced in NADP-ME and PCK than in NAD-ME grasses. Grasses grown in the field and in the glasshouse showed a similar pattern. Thus, caution should be exercised when using C4 plants under varying environmental conditions to monitor the concentration or carbon isotopic composition of atmospheric CO2 in field/glasshouse studies or climate reconstruction.